Photosynthesis, stomatal conductance and terpene emission response to water availability in dry and mesic Mediterranean forests

Water stress results in a reduction of the metabolism of plants and in a reorganization of their use of resources geared to survival. In the Mediterranean region, periods of drought accompanied by high temperatures and high irradiance occur in summer. Plants have developed various mechanisms to survive in these conditions by resisting, tolerating or preventing stress. We used three typical Mediterranean tree species in Israel, Pinus halepensis L., Quercus calliprinos and Quercus ithaburensis Webb, as models for studying some of these adaptive mechanisms. We measured their photosynthetic rates (A), stomatal conductance (g s), and terpene emission rates during spring and summer in a geophysical gradient from extremely dry to mesic from Yatir (south, arid) to Birya (north, moist) with intermediate conditions in Solelim. A and g s of P. halepensis were threefold higher in Birya than in Yatir where they remained very low both seasons. Quercus species presented 2-3-fold higher A and g s but with much more variability between seasons, especially for Q. ithaburensis with A and g s that decreased 10-30-fold from spring to summer. Terpene emission rates for pine were not different regionally in spring but they were 5-8-fold higher in Birya than in Yatir in summer (P < 0.05). Higher emissions were also observed in Solelim for the drought resistant Q. ithaburensis (P < 0.001) but not for Q. calliprinos. α-Pinene followed by limonene and 3-carene were the dominant terpenes. Warmer summer conditions result in increased Terpene emission rates except under severe drought, in which case they strongly decreas

Vapour pressure deficit was not a primary limiting factor for gas exchange in an irrigated, mature dryland Aleppo pine forest

Climate change is often associated with increasing vapour pressure deficit (VPD) and changes in soil moisture (SM). While atmospheric and soil drying often co-occur, their differential effects on plant functioning and productivity remain uncertain. We investigated the divergent effects and underlying mechanisms of soil and atmospheric drought based on continuous, in situ measurements of branch gas exchange with automated chambers in a mature semiarid Aleppo pine forest. We investigated the response of control trees exposed to combined soil‒atmospheric drought (low SM, high VPD) during the rainless Mediterranean summer and that of trees experimentally unconstrained by soil dryness (high SM; using supplementary dry season water supply) but subjected to atmospheric drought (high VPD). During the seasonal dry period, branch conductance (g$_{br}$), transpiration rate (E) and net photosynthesis (A$_{net}$) decreased in low-SM trees but greatly increased in high-SM trees. The response of E and g$_{br}$ to the massive rise in VPD (to 7 kPa) was negative in low-SM trees and positive in high-SM trees. These observations were consistent with predictions based on a simple plant hydraulic model showing the importance of plant water potential in the g$_{br}$ and E response to VPD. These results demonstrate that avoiding drought on the supply side (SM) and relying on plant hydraulic regulation constrains the effects of atmospheric drought (VPD) as a stressor on canopy gas exchange in mature pine trees under field conditions

Assessing model performance via the most limiting environmental driver in two differently stressed pine stands

Climate change will impact forest productivity worldwide. Forecasting the magnitude of such impact, with multiple environmental stressors changing simultaneously, is only possible with the help of process-based models. In order to assess their performance, such models require careful evaluation against measurements. However, direct comparison of model outputs against observational data is often not reliable, as models may provide the right answers due to the wrong reasons. This would severely hinder forecasting abilities under unprecedented climate conditions. Here, we present a methodology for model assessment, which supplements the traditional output-to-observation model validation. It evaluates model performance through its ability to reproduce observed seasonal changes of the most limiting environmental driver (MLED) for a given process, here daily gross primary productivity (GPP). We analyzed seasonal changes of the MLED for GPP in two contrasting pine forests, the Mediterranean Pinus halepensis Mill. Yatir (Israel) and the boreal Pinus sylvestris L. Hyytiala (Finland) from three years of eddy-covariance flux data. Then, we simulated the same period with a state-of-the-art process-based simulation model (LandscapeDNDC). Finally, we assessed if the model was able to reproduce both GPP observations and MLED seasonality. We found that the model reproduced the seasonality of GPP in both stands, but it was slightly overestimated without site-specific fine-tuning. Interestingly, although LandscapeDNDC properly captured the main MLED in Hyytiala (temperature) and in Yatir (soil water availability), it failed to reproduce high-temperature and high-vapor pressure limitations of GPP in Yatir during spring and summer. We deduced that the most likely reason for this divergence is an incomplete description of stomatal behavior. In summary, this study validates the MLED approach as a model evaluation tool, and opens up new possibilities for model improvement.Peer reviewe

Potential and limitations of inferring ecosystem photosynthetic capacity from leaf functional traits

The aim of this study was to systematically analyze the potential and limitations of using plant functional trait observations from global databases versus in situ data to improve our understanding of vegetation impacts on ecosystem functional properties (EFPs). Using ecosystem photosynthetic capacity as an example, we first provide an objective approach to derive robust EFP estimates from gross primary productivity (GPP) obtained from eddy covariance flux measurements. Second, we investigate the impact of synchronizing EFPs and plant functional traits in time and space to evaluate their relationships, and the extent to which we can benefit from global plant trait databases to explain the variability of ecosystem photosynthetic capacity. Finally, we identify a set of plant functional traits controlling ecosystem photosynthetic capacity at selected sites. Suitable estimates of the ecosystem photosynthetic capacity can be derived from light response curve of GPP responding to radiation (photosynthetically active radiation or absorbed photosynthetically active radiation). Although the effect of climate is minimized in these calculations, the estimates indicate substantial interannual variation of the photosynthetic capacity, even after removing site-years with confounding factors like disturbance such as fire events. The relationships between foliar nitrogen concentration and ecosystem photosynthetic capacity are tighter when both of the measurements are synchronized in space and time. When using multiple plant traits simultaneously as predictors for ecosystem photosynthetic capacity variation, the combination of leaf carbon to nitrogen ratio with leaf phosphorus content explains the variance of ecosystem photosynthetic capacity best (adjusted R-2 = 0.55). Overall, this study provides an objective approach to identify links between leaf level traits and canopy level processes and highlights the relevance of the dynamic nature of ecosystems. Synchronizing measurements of eddy covariance fluxes and plant traits in time and space is shown to be highly relevant to better understand the importance of intra-and interspecific trait variation on ecosystem functioning.Peer reviewe

Resilience to seasonal heat wave episodes in a Mediterranean pine forest

Short-term, intense heat waves (hamsins) are common in the eastern Mediterranean region and provide an opportunity to study the resilience of forests to such events that are predicted to increase in frequency and intensity. * The response of a 50-yr-old Aleppo pine (Pinus halepensis) forest to hamsin events lasting 1–7 d was studied using 10 yr of eddy covariance and sap flow measurements. * The highest frequency of heat waves was c. four per month, coinciding with the peak productivity period (March–April). During these events, net ecosystem carbon exchange (NEE) and canopy conductance (gc) decreased by c. 60%, but evapotranspiration (ET) showed little change. Fast recovery was also observed with fluxes reaching pre-stress values within a day following the event. NEE and gc showed a strong response to vapor pressure deficit that weakened as soil moisture decreased, while sap flow was primarily responding to changes in soil moisture. On an annual scale, heat waves reduced NEE and gross primary productivity by c. 15% and 4%, respectively. * Forest resilience to short-term extreme events such as heat waves is probably a key to its survival and must be accounted for to better predict the increasing impact on productivity and survival of such events in future climates

Assessing climatic benefits from forestation potential in semi-arid lands

Forestation actions are a major tool for both climate-change mitigation and biodiversity conservation. We address two weaknesses in this approach: the little attention given to the negative effects of reduced albedo associated with forestation in many regions, and ignoring the potential of drylands that account for 40% of the global potential land area for forestation. We propose an approach to identify suitable land for forestation and quantify its ‘net equivalent carbon stock change’ over 80 years of forest lifetime (NESC), accounting for both carbon sequestration and albedo changes. We combined remote-sensing tools with data-based estimates of surface parameters and with published climate matrices, to identify suitable land for forestation actions. We then calculated the cumulative (over 80 years) ‘net sequestration potential’ (ΔSP), the ‘emission equivalent of shortwave radiation forcing’ (EESF) due to changes in surface albedo, and, in turn, the combined NESC = ΔSP−EESF, of planting forests with >40% tree-cover. Demonstrating our approach in a large climatically diverse state (Queensland), we identified 14.5 million hectares of potential forestation land in its semi-arid land and show that accounting for the EESF, reduces the climatic benefits of the ΔSP by almost 50%. Nevertheless, it results in a total NESC of 0.72 Gt C accumulated by the end of the century, and 80 years of forestation cycle. This estimated NESC is equivalent to 15% of the projected carbon emissions for the same period in Queensland, for a scenario of no change in emission rates during that period. Our approach extends restoration efforts by identifying new land for forestation and carbon sequestration but also demonstrates the importance of quantifying the climatic value of forestation in drylands

Large variations in afforestation-related climate cooling and warming effects across short distances

The time for the climatic benefits of afforestation via carbon-sequestration to offset the warming effects of reduced albedo and longwave radiation emission varies greatly across short distances, according to a study of paired forested and non-forested ecosystems along an aridity gradient

Differential ecophysiological response of a major Mediterranean pine species to climatic gradient

The rate of migration and in situ genetic variation in forest trees may not be sufficient to compete with the current rapid rate of climate change. Ecophysiological adjustments of key traits, however, could complement these processes and allow sustained survival and growth across a wide range of climatic conditions. This was tested in Pinus halepensis Miller by examining seven physiological and phenological parameters in five provenances growing in three common garden plots along a climatic transect from meso-Mediterranean (MM) to thermo-Mediterranean (TM) and semi-arid (SA) climates. Differential responses to variations in ambient climatic conditions were observed in three key traits: (i) growing season length decreased with drying in all provenances examined (from 165 under TM climate to 100 days under SA climate, on average); (ii) water use efficiency (WUE) increased with drying, but to a different extent in different provenances, and on average from 80, to 95, to 110 µmol CO(2) mol(-1) H(2)O under MM, TM and SA climates, respectively; (iii) xylem native embolism was stable across climates, but varied markedly among different provenances (percent loss of conductivity, was below 5% in two provenances and above 35% in others). The results indicated that changes in growing season length and WUE were important contributors to tree growth across climates, whereas xylem native embolism negatively correlated with tree survival. The results indicated that irrespective of slow processes (e.g., migration, genetic adaptation), the capacity for ecophysiological adjustments combined with existing variations among provenances could help sustain P. halepensis, a major Mediterranean tree species, under relatively extreme warming and drying climatic trends

Disentangling Soil, Shade, and Tree Canopy Contributions to Mixed Satellite Vegetation Indices in a Sparse Dry Forest

Remote sensing (RS) for vegetation monitoring can involve mixed pixels with contributions from vegetation and background surfaces, causing biases in signals and their interpretations, especially in low-density forests. In a case study in the semi-arid Yatir forest in Israel, we observed a mismatch between satellite (Landsat 8 surface product) and tower-based (Skye sensor) multispectral data and contrasting seasonal cycles in near-infrared (NIR) reflectance. We tested the hypothesis that this mismatch was due to the different fractional contributions of the various surface components and their unique reflectance. Employing an unmanned aerial vehicle (UAV), we obtained high-resolution multispectral images over selected forest plots and estimated the fraction, reflectance, and seasonal cycle of the three main surface components (canopy, shade, and sunlit soil). We determined that the Landsat 8 data were dominated by soil signals (70%), while the tower-based data were dominated by canopy signals (95%). We then developed a procedure to resolve the canopy (i.e., tree foliage) normalized difference vegetation index (NDVI) from the mixed satellite data. The retrieved and corrected canopy-only data resolved the original mismatch and indicated that the spatial variations in Landsat 8 NDVI were due to differences in stand density, while the canopy-only NDVI was spatially uniform, providing confidence in the local flux tower measurements